Consequences of Extended Egg Retention in the Eastern Fence Lizard (Sceloporus undulatus)

Size: px
Start display at page:

Download "Consequences of Extended Egg Retention in the Eastern Fence Lizard (Sceloporus undulatus)"

Transcription

1 Journal of Herpetology, Vol. 37, No. 2, pp , 2003 Copyright 2003 Society for the Study of Amphibians and Reptiles Consequences of Extended Egg Retention in the Eastern Fence Lizard (Sceloporus undulatus) DANIEL A. WARNER 1 AND ROBIN M. ANDREWS Department of Biology, Virginia Polytechnic Institute and State University, Blacksburg, Virginia 24061, USA ABSTRACT. Egg retention beyond the normal time of oviposition occurs frequently in oviparous squamate reptiles and is thought to be a response to unfavorable nesting conditions. During studies of the Eastern Fence Lizard (Sceloporus undulatus), we obtained data on the effects of extended egg retention on embryonic development, hatchling phenotypes, and posthatching survival under natural field conditions. Females that retained eggs beyond the normal time of oviposition produced heavier eggs with embryos more advanced (by one stage unit) at the time of oviposition than females that did not retain eggs for extended periods. Egg retention did not affect any hatchling phenotype (i.e., body size, thermal preference, running speed, desiccation rate, growth rate) but had a significant positive effect on posthatching survival in the field. However, the mechanism by which extended egg retention affects posthatching survival remains unclear. Our results have implications for the evolution of viviparity, but carefully designed experiments are needed to further understand the causes and consequences of extended egg retention. The retention of eggs in the oviduct beyond the normal time of oviposition is common in squamate reptiles and may occur in response to dry environmental conditions (Jones et al., 1991; Andrews and Rose, 1994; Radder et al., 1998). For example, if nest sites are too dry, females may delay nesting and either retain eggs until conditions become favorable or oviposit (without nesting) when egg retention is no longer physiologically possible (Mathies and Andrews, 1996; Warner and Andrews, 2002a). The developmental consequences of extended egg retention vary among species. For instance, extended egg retention results in developmental arrest for embryos of Urosaurus ornatus and Sceloporus undulatus, but embryogenesis continues in utero to a stage that is near the time of hatching in Sceloporus scalaris (Andrews and Mathies, 2000). For species in which development is arrested, embryogenesis is resumed after eggs are laid. Moreover, during the time embryos are retained in utero, the maternal embryonic environment can affect the hydration and size of hatchlings (Andrews and Mathies, 2000). Extended egg retention may thus affect offspring phenotype or fitness, but the long-term consequences of extended egg retention are unknown. We evaluated effects of extended egg retention on embryonic development, hatchling phenotypes, and posthatching survival of the Eastern Fence Lizard (Sceloporus undulatus), a species that oviposits when embryos are at stages on the Dufaure and Hubert (1961) scale. The 1 Corresponding Author. Present address: School of Biological Sciences, The University of Sydney, Sydney, New South Wales 2006, Australia. results presented in this paper were obtained while conducting studies on the nesting behavior of S. undulatus (Warner and Andrews, 2002a) and on the associations between phenotypes and survival of hatchling S. undulatus in the field (Warner, 2001; Warner and Andrews, 2002b). MATERIALS AND METHODS Gravid S. undulatus (N 23) were collected between 15 May and 19 June 1999 in Jefferson National Forest (Montgomery County) near Blacksburg, Virginia. Females were brought to facilities at Virginia Polytechnic Institute and State University where they were housed in a m enclosure. This enclosure was specifically designed to study nesting behavior of S. undulatus (Warner and Andrews, 2002a). The enclosure provided areas for basking and nine plastic containers ( cm), filled with a mixture of vermiculite and peat moss (1:1 ratio), provided nesting substrate. All lizards readily had access to the nine nesting containers which where placed adjacent to one another in a3 3 array. Thus, the total area available for nesting was cm. The enclosure was illuminated by heat lamps and two 1.3-m Vitalites. Females were placed in the enclosure as they were captured and removed from the enclosure after they oviposited. Thus, no more than 10 females were housed in the enclosure at a given time. Females were fed crickets and wax worm larvae dusted with a vitamin-mineral mix, and watered daily. See Warner and Andrews (2002a) for a complete description of the enclosure. Seventeen females constructed nests and exhibited nest construction behavior similar to that of females in the wild (Warner and Andrews, 2002a). Six females oviposited on the

2 310 D. A. WARNER AND R. M. ANDREWS surface of the substrate (without nesting). Eggs were either removed from nests or removed from the enclosure (eggs from females that did not nest), and one egg from each of the 23 clutches used in this study was sampled to determine the stage of the embryo at oviposition (staged according to Dufaure and Hubert, 1961). One embryo provides a reliable representation of an entire clutch because embryonic stage at oviposition does not vary within clutches (DeMarco, 1992). We could not determine the embryonic stage of one clutch from a female that nested because we did not find the embryo in the egg sampled. Immediately after oviposition, all remaining eggs were weighed and placed individually in glass jars (65 ml) containing vermiculite and covered with clear plastic wrap sealed with a rubberband. Jars were placed in an environmental chamber set at a constant 28 C and rotated to different positions within the chamber to minimize possible effects of temperature gradients within the chamber (Warner and Andrews, 2002b). After hatching, we marked each hatchling individually by unique toe-clips and measured several phenotypic traits of each individual. For each individual, we measured snout vent length (SVL), tail length (TL), and mass, and body shape was calculated as mass 0.3 /SVL. We also measured thermal preference, locomotor performance, desiccation rate, and growth rate for each hatchling in the laboratory. Thermal preference was measured by placing lizards within a thermal gradient (at 1 2 days of age). The thermal gradient contained nine thermal couples spaced at 10-cm intervals that measured air temperatures along the length (80 cm) of the gradient. Air temperatures varied linearly from C along the gradient. The air temperature at the position of the lizard within the gradient reflected the lizard s body temperature. The position of the lizard within the gradient was recorded every 10 min for an hour and air temperatures within the gradient were recorded at the beginning, middle, and end of the hour. Lizard position was then converted to body temperature by regression analysis. Selected body temperature was calculated as the mean of the six temperature measurements over the 1-h observation period. See Qualls and Andrews (1999) and Warner and Andrews (2002b) for complete descriptions of protocols. Locomotor performance was measured by chasing hatchlings along a 1-m electronically timed racetrack (at 3 4 days of age). The racetrack consisted of five infrared photocells (spaced at 25-cm intervals) connected to an electronic stopwatch. Hatchlings were placed at the beginning of the racetrack and gently prodded with a paintbrush if they did not run or stopped. Locomotor performance over 1 m was measured three times for each individual and was assessed as the fastest speed (m/s) over 25 cm and 1 m. All running trials were conducted inside a walk-in environmental chamber set at 30 C. See Warner and Andrews (2002b) for a complete description of protocols. Desiccation rate was measured by placing lizards in a desiccator set at 0% relative humidity and at a constant 30 C (at 6 7 days of age). Body mass was recorded before hatchlings were placed in the desiccator and again after 2 h. Desiccation rate was assessed as short-term evaporative water loss (change in body mass/ h). Growth in both SVL and mass was calculated as a size specific growth rate: the difference between an individual s natural log-transformed SVL or mass at the time of release and hatching divided by the number of days between measurements. Each hatchling was subsequently released (at an average age of nine days) between 24 July and 8 September The release site was located in Jefferson National Forest near Blacksburg, Virginia and included a 1500-m 2 forest clearing, a smaller clearing (500 m 2 ), and an abandoned dirt road extending 80 m from the site. The open areas contained large woody debris and scattered small shrubs, which provided habitat for S. undulatus. The entire site was surrounded by dense forest, which provided a natural boundary because hatchling S. undulatus do not disperse through the forest (see Warner and Andrews, 2002b). We searched the site for hatchlings about twice weekly from 31 July to 5 December 1999 and weekly from 5 March to 6 July The perimeter of the field site was searched thoroughly and few (6% of those released) hatchlings were found in the surrounding forest. Thus, dispersal did not appear to bias our recapture success, and disappearances from the site were likely caused by death. Furthermore, our recapture rates were nearly identical to recapture rates of S. undulatus hatchlings released in an enclosed area where dispersal was not possible (Niewiarowski and Roosenburg, 1993). Of 220 hatchlings released, 130 were recaptured (by hand) at least once after release. After recapture, hatchlings were identified by their toe clip, weighed, measured (SVL and TL) and then released where they were captured. Thus, this mark-recapture study allowed us to measure survival under natural field conditions. Detailed descriptions of the above protocols are presented by Warner and Andrews (2002b). To determine the effect of prolonged egg retention on hatchling phenotypes and survival, we compared clutches from females that nested (N 17) with clutches from females that did

3 EGG RETENTION IN SCELOPORUS UNDULATUS 311 TABLE 1. Comparisons of maternal characteristics, egg characteristics, and hatchling phenotypes between females that nested and females that did not nest. Significant P-values are in bold type. Statistical tests were performed with ANOVA except where noted. Days females were in captivity Embryonic stage at oviposition Clutch size Clutch mass (g) Relative clutch mass (g) 1 Egg mass at oviposition (g) 1 Incubation period (days) 2 Hatching success (%) Date of hatching (Julian day) Hatchling SVL (mm) 3 Hatchling mass (g) 3 Tail length (mm) 4 Body shape (mass 0.3 /SVL) Thermal preference ( C) Running speed over 25 cm (m/s) 4 Running speed over 1 m (m/s) 4 Desiccation rate ( g/h) 5 Growth in SVL (log mm/day) Growth in mass (log g/day) Date of release (Julian day) Females that nested Mean 1SE Females that did not nest Mean 1 SE Statistical test F 1,21 8.0, P F 1, , P F 1,21 1.1, P F 1,21 0.0, P F 1,20 2.0, P F 1,20 5.0, P F 1,19 1.1, P F 1,21 0.1, P F 1,21 4.6, P F 1,20 0.2, P F 1,20 3.8, P F 1,19 0.1, P F 1,21 1.4, P F 1,21 0.5, P F 1,20 0.0, P F 1,20 0.0, P F 1,20 0.7, P F 1,21 0.0, P F 1,21 1.3, P F 1,21 5.4, P Superscripts denote traits analyzed with ANCOVA using the following covariates: 1 female mass after oviposition, 2 embryonic stage, 3 egg mass at oviposition, 4 snout-vent length at hatching, 5 body mass at hatching. Least-squares means are reported. not nest but oviposited on the surface of the substrate (N 6). We made these comparisons assuming that the females that oviposited on the substrate surface had retained their eggs beyond the normal time of oviposition (see Results). This assumption was based on the observation that Sceloporus lizards retain eggs beyond the normal time of oviposition and oviposit on the substrate surface if suitable nesting conditions do not become available (Jones et al., 1991; Andrews and Rose, 1994; Mathies and Andrews, 1996). The six females that oviposited on the surface may have perceived nesting conditions as unsuitable. Analyses of variance (ANOVA) and covariance (ANCOVA) were used to evaluate the effect of egg retention on embryonic development, hatchling phenotypes, and survival in the field. Female body mass was used as a covariate when analyzing the effect of egg retention on egg mass and clutch mass. Embryonic stage at oviposition was used as a covariate when analyzing incubation period. Egg mass was used as a covariate when analyzing hatchling body size (SVL and mass). Hatchling SVL and body mass were used as covariates when analyzing hatchling running speed and desiccation rate, respectively. All analyses of hatchling phenotypes were based on clutch means for each trait. Hatchling survival was analyzed at three time periods: survival at six and 12 weeks after release and again the following March If hatchlings were not recaptured, they were assumed dead (see above). Because egg retention by females that did not nest caused a delay in hatching date and subsequent delay in release of hatchlings (see Results), we performed an additional analysis to separate effects of release date and egg retention on hatchling survival. The release date of hatchlings from females that nested was distributed over a broad time range and, thus, allowed us to divide their clutches into early (19 July to 8 August) and late (18 August to 7 September) release time periods. Survival was then compared (ANOVA) at six weeks, 12 weeks, and March between clutches with hatchlings released early and clutches with hatchlings released late. Survival analyses were based on the percentage of individuals recaptured for each clutch. RESULTS Egg retention time, embryonic stage, and egg mass at oviposition differed significantly between females that nested and those that did not nest (Table 1). Females that did not nest were in captivity for an average of 12 days longer than females that nested. Moreover, females that did not nest oviposited when embryos were one stage more advanced than females that

4 312 D. A. WARNER AND R. M. ANDREWS FIG. 1. Relationship between the number of days females were in captivity and the stage of their embryos at oviposition (N 22, r , P 0.003). The numbers of datapoints that overlap are indicated on the graph. nested. The number of days females were held in captivity was positively related to embryonic stage at oviposition (r , P 0.003; Fig. 1). Females that did not nest produced heavier eggs at the time of oviposition than females that nested. However, clutch size, clutch mass, incubation period, and hatching success did not differ between females that nested and those that did not nest (Table 1). Delayed oviposition by females that retained eggs resulted in a shift in hatching dates by an average of 11 days (Table 1, Fig. 2). These observations clearly show that females that did not nest retained eggs beyond their normal time of oviposition. Egg retention had no effect on hatchling body size (mass or SVL; Table 1), despite the significant effect on egg size, even when body size was not corrected for egg size (mass: F 1,21 0.3, P 0.586; SVL: F 1,21 0.1, P 0.791). Moreover, egg retention had no effect on any other morphological (tail length and body shape) or performance traits (thermal preference, running speed, desiccation rate, and growth) of the hatchlings (Table 1). However, females that retained their eggs produced clutches with greater posthatching survival in the field than females that nested normally (Fig. 3). This pattern of survival was highly significant at 12 weeks after release and in the following March Hatchling release date did not influence posthatching survival in the field. In analyses using only clutches from females that nested, hatchlings released at the early and late time periods did not differ in survival at six weeks after release (F 1,12 0.0, P 0.980), 12 weeks after re- FIG. 2. Relationship between mean oviposition date and mean hatching date for clutches from females that nested (open circles) and females that did not nest (solid circles; r , P 0.001). FIG. 3. Comparison of mean posthatching survival of clutches from females that nested (hatched bars) and females that retained eggs for prolonged periods (open bars). Mean survival is from release to six weeks, from release to 12 weeks, and from release to the following March Bars represent 1 SE.

5 EGG RETENTION IN SCELOPORUS UNDULATUS 313 lease (F 1,12 0.6, P 0.471) and at the following March 2000 (F 1,12 2.3, P 0.154). DISCUSSION Differences in oviposition dates between females that nested and females that oviposited on the substrate were caused by egg retention beyond the normal time of oviposition by the females that did not nest. Females that did not nest were in captivity 12 days longer (prior to oviposition) than females that nested. Moreover, females that oviposited on the substrate did so when embryos were at a slightly more advanced embryonic stage than embryos from females that nested. This result parallels that of another study on S. undulatus in which females exposed to unsuitable nesting conditions retained their eggs an average of 10 days beyond the normal time of oviposition (Andrews and Mathies, 2000). During that 10-day period, control embryos (from females that nested) reached stage 32, whereas retained embryos reached stage 30; the rate of embryonic development is retarded during egg retention (Andrews and Mathies, 2000). Egg retention would thus not be obvious from the stage of embryos alone in our study or theirs. Females that retained eggs produced eggs that were 9% heavier than those produced by females that nested. The relatively large egg mass of females that retained eggs did not result in relatively large hatchlings, suggesting the increased egg mass was caused by water uptake during the 12-day period of prolonged egg retention. Reptile eggs must take up water after oviposition for successful development, and oviposited eggs of S. undulatus double or triple in mass before hatching (Warner and Andrews, 2002b). Water uptake in utero, however, may be restricted because of physical constraints within the female s oviducts (Mathies and Andrews, 1996). Indeed, egg expansion within the oviduct may eventually cause the female to oviposit even if no suitable nesting sites are available. Egg retention did not affect the morphology or performance of hatchlings, but it did increase their survival after release in the field. The mechanism by which egg retention increased the survival of hatchlings, however, cannot be identified. We provide three possible explanations why egg retention increased the survival of hatchlings. First, release date could influence hatchling survival (Ferguson and Bohlen, 1978; Sinervo and Doughty, 1996; Madsen and Shine, 1998). In our study, the 11-day delay in hatching date (and subsequent delay in release dates) means that hatchlings from eggs that had been retained were released 12 days later than hatchlings from eggs that had not been retained. Release date, however, was not related to hatchling survival in the field, possibly because release dates of hatchlings that were from retained clutches and nested clutches were separated by a relatively short time period (only 12 days). Our results are in contrast to a previous study on the same population, which found that individuals that hatched late had higher survival than those that hatched early (Andrews et al., 2000). Release dates in that study, however, extended over a two-month period. The explanation of Andrews et al. (2000) was that individuals that hatched late were not exposed to predators for as long a period as those that hatched early. Second, egg retention did not influence any hatchling phenotype that we measured and, therefore, did not indirectly influence survival through these phenotypes. Perhaps, however, egg retention influenced survival through its effect on other unmeasured phenotypes, whether morphological, physiological, or behavioral. Third, the effect of egg retention on survival may be an artifact because of confounding effects of clutch and egg retention. Clutch has strong effects on posthatching survival in reptiles (Madsen and Shine, 1998; Warner and Andrews, 2002b). Thus, clutch (maternal) effects other than egg retention per se may explain the difference in survival between clutches that were retained in utero for an extended period and those that were laid at the normal time of oviposition. Extended periods of egg retention have important implications for the evolution of viviparity. Viviparity is thought to have evolved through an increased capacity to retain eggs coupled with the maintenance of embryogenesis in utero (Andrews and Mathies, 2000). Our data suggest that extended egg retention could provide a survival advantage for offspring, but the mechanism by which extended egg retention affects survival remains unclear. An important point to consider, however, is that egg retention is beneficial for oviparous females only if females eventually encounter suitable nesting substrates. In nature, eggs laid on the surface would not survive, but given the variety of microenvironments in the field compared to those provided in the laboratory, females are likely to eventually encounter a suitable nesting site during a period of prolonged egg retention. Furthermore, the lack of nesting behavior by females in the laboratory probably rarely occurs in nature, if it does at all. Nevertheless, the positive affect of extended egg retention on survival may be an important step that favors the evolution of viviparity and could precede the ability to retain eggs in utero for prolonged periods. Ad-

6 314 D. A. WARNER AND R. M. ANDREWS ditional studies that are specifically designed to evaluate long-term effects of extended egg retention are needed. Furthermore, comparative studies of closely related taxa that vary in their capacity to retain eggs will help to identify causes and consequences of extended egg retention. Acknowledgments. Funding was provided by grants from Sigma Xi, the Chicago Herpetological Society, and the Graduate Student Assembly of Virginia Polytechnic Institute and State University. We thank C. Astor, M. Lovern, R. McCleary, K. Passek, and J. Thomson for their assistance in the field and laboratory. All lizards were collected under permit of the Virginia Department of Game and Inland Fisheries. This project was approved by the Virginia Polytechnic Institute and State University Animal Care Committee (proposal BIOL). LITERATURE CITED ANDREWS, R. M., AND T. MATHIES Natural history of reptilian development: constraints on the evolution of viviparity. Bioscience 50: ANDREWS, R. M., AND B. R. ROSE Evolution of viviparity: constraints of egg retention. Physiological Zoology 67: ANDREWS, R. M., T. MATHIES, AND D. A. WARNER Effect of incubation temperature on morphology, growth, and survival of juvenile Sceloporus undulatus. Herpetological Monographs 14: DE MARCO, V Embryonic development times and egg retention in four species of sceloporine lizards. Functional Ecology 6: DUFAURE, J. P., AND J. HUBERT Table de développement du lézard vivipare: Lacerta (Zootoca) vivipara Jacquin. Archives Anatomie Microscopie Morphologie Expérimental 50: FERGUSON, G. W., AND C. H. BOHLEN Demographic analysis: a tool for the study of natural selection of behavioral traits. In N. Greenberg and P. D. MacLean (eds.), Behavior and Neurology of Lizards, pp Department of Health, Education, and Welfare Publication (ADM) , Washington, DC. JONES, R. E., C. R. PROPPER, M.S.RAND, AND H. B. AUSTIN Loss of nesting behavior and the evolution of viviparity in reptiles. Ethology 88: MADSEN, T.,AND R. SHINE Quantity or quality? Determinants of maternal reproductive success in tropical pythons (Liasis fuscus). Proceedings of the Royal Society of London, B Biological Sciences 265: MATHIES, T., AND R. M. ANDREWS Extended egg retention and its influence on embryonic development and egg water balance: implications for the evolution of viviparity. Physological Zoology 69: NIEWIAROWSKI, P. H., AND W. ROOSENBURG Reciprocal transplant reveals sources of variation in growth rates of the lizard Sceloporus undulatus. Ecology 74: QUALLS, C.P.,AND R. M. ANDREWS Cold climates and the evolution of viviparity in reptiles: cold incubation temperatures produce poorquality offspring in the lizard, Sceloporus virgatus. Biological Journal of the Linnean Society 67: RADDER, R. S., B. A. SHANBHAG, AND S. K. SAIDAPUR Prolonged oviductal egg retention arrests embryonic growth at stage 34 in captive Calotes versicolor. Herpetological Review 29: SINERVO, B., AND P. DOUGHTY The interactive effects of offspring size and timing of reproduction on offspring reproduction; experimental, maternal, and quantitative genetic aspects. Evolution 50: WARNER, D. A Phenotypes and survival of hatchling lizards. Unpubl. master s thesis, Virginia Polytechnic Institute and State Univ., Blacksburg. WARNER, D. A., AND R. M. ANDREWS. 2002a. Nestsite selection in relation to temperature and moisture by the lizard Sceloporus undulatus. Herpetologica 58: b. Laboratory and field experiments identify sources of variation in phenotypes and survival of hatchling lizards. Biological Journal of the Linnean Society 76: Accepted: 6 December 2002.

Incubation temperature and phenotypic traits of Sceloporus undulatus: implications for the northern limits of distribution

Incubation temperature and phenotypic traits of Sceloporus undulatus: implications for the northern limits of distribution DOI 10.1007/s00442-006-0583-0 ECOPHYSIOLOGY Incubation temperature and phenotypic traits of Sceloporus undulatus: implications for the northern limits of distribution Scott L. Parker Æ Robin M. Andrews

More information

PHYSIOLOGICAL AND ECOLOGICAL CONSTRAINTS ON THE EVOLUTION OF VIVIPARITY IN SCELOPORINE LIZARDS. Scott L. Parker

PHYSIOLOGICAL AND ECOLOGICAL CONSTRAINTS ON THE EVOLUTION OF VIVIPARITY IN SCELOPORINE LIZARDS. Scott L. Parker PHYSIOLOGICAL AND ECOLOGICAL CONSTRAINTS ON THE EVOLUTION OF VIVIPARITY IN SCELOPORINE LIZARDS Scott L. Parker Dissertation submitted to the faculty of Virginia Polytechnic Institute and State University

More information

Seasonal Shifts in Reproductive Investment of Female Northern Grass Lizards ( Takydromus septentrionalis

Seasonal Shifts in Reproductive Investment of Female Northern Grass Lizards ( Takydromus septentrionalis Seasonal Shifts in Reproductive Investment of Female Northern Grass Lizards (Takydromus septentrionalis) from a Field Population on Beiji Island, China Author(s): Wei-Guo Du and Lu Shou Source: Journal

More information

PHENOTYPES AND SURVIVAL OF HATCHLING LIZARDS. Daniel A. Warner. MASTER OF SCIENCE in Biology

PHENOTYPES AND SURVIVAL OF HATCHLING LIZARDS. Daniel A. Warner. MASTER OF SCIENCE in Biology PHENOTYPES AND SURVIVAL OF HATCHLING LIZARDS Daniel A. Warner Thesis submitted to the Faculty of Virginia Polytechnic Institute and State University in partial fulfillment of the requirements for the degree

More information

Embryonic responses to variation in oviductal oxygen in the lizard Sceloporus undulatus from New Jersey and South Carolina, USA

Embryonic responses to variation in oviductal oxygen in the lizard Sceloporus undulatus from New Jersey and South Carolina, USA Blackwell Science, LtdOxford, UKBIJBiological Journal of the Linnean Society0024-4066The Linnean Society of London, 2004? 2004 83? 289299 Original Article Biological Journal of the Linnean Society, 2004,

More information

Geographic variation in lizard phenotypes: importance of the incubation environment

Geographic variation in lizard phenotypes: importance of the incubation environment Biological Journal of the Linnean Society (1998), 64: 477 491. With 3 figures Article ID: bj980236 Geographic variation in lizard phenotypes: importance of the incubation environment FIONA J. QUALLS AND

More information

FEMALE PHENOTYPE, LIFE HISTORY, AND REPRODUCTIVE SUCCESS IN FREE-RANGING SNAKES (TROPIDONOPHIS MAIRII)

FEMALE PHENOTYPE, LIFE HISTORY, AND REPRODUCTIVE SUCCESS IN FREE-RANGING SNAKES (TROPIDONOPHIS MAIRII) Ecology, 86(10), 2005, pp. 2763 2770 2005 by the Ecological Society of America FEMALE PHENOTYPE, LIFE HISTORY, AND REPRODUCTIVE SUCCESS IN FREE-RANGING SNAKES (TROPIDONOPHIS MAIRII) G. P. BROWN AND R.

More information

MATERNAL NEST-SITE CHOICE AND OFFSPRING FITNESS IN A TROPICAL SNAKE (TROPIDONOPHIS MAIRII, COLUBRIDAE)

MATERNAL NEST-SITE CHOICE AND OFFSPRING FITNESS IN A TROPICAL SNAKE (TROPIDONOPHIS MAIRII, COLUBRIDAE) Ecology, 85(6), 2004, pp. 1627 1634 2004 by the Ecological Society of America MATERNAL NEST-SITE CHOICE AND OFFSPRING FITNESS IN A TROPICAL SNAKE (TROPIDONOPHIS MAIRII, COLUBRIDAE) G. P. BROWN AND R. SHINE

More information

Effects of nest temperature and moisture on phenotypic traits of hatchling snakes (Tropidonophis mairii, Colubridae) from tropical Australia

Effects of nest temperature and moisture on phenotypic traits of hatchling snakes (Tropidonophis mairii, Colubridae) from tropical Australia Blackwell Publishing LtdOxford, UKBIJBiological Journal of the Linnean Society24-466The Linnean Society of London, 26? 26 891 159168 Original Article INCUBATION EFFECTS IN A SNAKE G. P. BROWN and R. SHINE

More information

Reproductive modes in lizards: measuring fitness. consequences of the duration of uterine retention of eggs

Reproductive modes in lizards: measuring fitness. consequences of the duration of uterine retention of eggs Functional Ecology 2008, 22, 332 339 doi: 10.1111/j.1365-2435.2007.01380.x Reproductive modes in lizards: measuring fitness Blackwell Publishing Ltd consequences of the duration of uterine retention of

More information

Egg environments have large effects on embryonic development, but have minimal consequences for hatchling phenotypes in an invasive lizard

Egg environments have large effects on embryonic development, but have minimal consequences for hatchling phenotypes in an invasive lizard 25..41 Biological Journal of the Linnean Society, 2012, 105, 25 41. With 6 figures Egg environments have large effects on embryonic development, but have minimal consequences for hatchling phenotypes in

More information

Effects of Incubation Temperature on Growth and Performance of the Veiled Chameleon (Chamaeleo calyptratus)

Effects of Incubation Temperature on Growth and Performance of the Veiled Chameleon (Chamaeleo calyptratus) JOURNAL OF EXPERIMENTAL ZOOLOGY 309A:435 446 (2008) A Journal of Integrative Biology Effects of Incubation Temperature on Growth and Performance of the Veiled Chameleon (Chamaeleo calyptratus) ROBIN M.

More information

Cold climates and the evolution of viviparity. produce poor-quality offspring in the lizard, in reptiles: cold incubation temperatures

Cold climates and the evolution of viviparity. produce poor-quality offspring in the lizard, in reptiles: cold incubation temperatures BiologicalJoumal of the Linriean Socieiv (l999), 67: 353-376. With 4 figures Article ID: bijl. 1998.0307, available online at http://~.idealit,rary.com on ID E bl 8 c Cold climates and the evolution of

More information

DOES VIVIPARITY EVOLVE IN COLD CLIMATE REPTILES BECAUSE PREGNANT FEMALES MAINTAIN STABLE (NOT HIGH) BODY TEMPERATURES?

DOES VIVIPARITY EVOLVE IN COLD CLIMATE REPTILES BECAUSE PREGNANT FEMALES MAINTAIN STABLE (NOT HIGH) BODY TEMPERATURES? Evolution, 58(8), 2004, pp. 1809 1818 DOES VIVIPARITY EVOLVE IN COLD CLIMATE REPTILES BECAUSE PREGNANT FEMALES MAINTAIN STABLE (NOT HIGH) BODY TEMPERATURES? RICHARD SHINE School of Biological Sciences,

More information

Thermal adaptation of maternal and embryonic phenotypes in a geographically widespread ectotherm

Thermal adaptation of maternal and embryonic phenotypes in a geographically widespread ectotherm International Congress Series 1275 (2004) 258 266 www.ics-elsevier.com Thermal adaptation of maternal and embryonic phenotypes in a geographically widespread ectotherm Michael J. Angilletta Jr. a, *, Christopher

More information

phenotypes of hatchling lizards, regardless of overall mean incubation temperature

phenotypes of hatchling lizards, regardless of overall mean incubation temperature Functional Ecology 2004 Seasonal shifts in nest temperature can modify the Blackwell Publishing, Ltd. phenotypes of hatchling lizards, regardless of overall mean incubation temperature R. SHINE* Biological

More information

ARTICLE IN PRESS. Zoology 113 (2010) 33 38

ARTICLE IN PRESS. Zoology 113 (2010) 33 38 Zoology 113 (2010) 33 38 Contents lists available at ScienceDirect Zoology journal homepage: www.elsevier.de/zool Effects of incubation temperature on hatchling phenotypes in an oviparous lizard with prolonged

More information

Phenotypic Effects of Thermal Mean and Fluctuations on Embryonic Development and Hatchling Traits in a Lacertid Lizard, Takydromus septentrionalis

Phenotypic Effects of Thermal Mean and Fluctuations on Embryonic Development and Hatchling Traits in a Lacertid Lizard, Takydromus septentrionalis JOURNAL OF EXPERIMENTAL ZOOLOGY 9A:138 146 (08) A Journal of Integrative Biology Phenotypic Effects of Thermal Mean and Fluctuations on Embryonic Development and Hatchling Traits in a Lacertid Lizard,

More information

Maternal Thermal Effects on Female Reproduction and Hatchling Phenotype in the Chinese Skink (Plestiodon chinensis)

Maternal Thermal Effects on Female Reproduction and Hatchling Phenotype in the Chinese Skink (Plestiodon chinensis) Asian Herpetological Research 2018, 9(4): 250 257 DOI: 10.16373/j.cnki.ahr.180056 ORIGINAL ARTICLE Maternal Thermal Effects on Female Reproduction and Hatchling Phenotype in the Chinese Skink (Plestiodon

More information

Is Parental Care the Key to Understanding Endothermy in Birds and Mammals?

Is Parental Care the Key to Understanding Endothermy in Birds and Mammals? vol. 162, no. 6 the american naturalist december 2003 Is Parental Care the Key to Understanding Endothermy in Birds and Mammals? Michael J. Angilletta, Jr., * and Michael W. Sears Department of Life Sciences,

More information

Influence of Incubation Temperature on Morphology, Locomotor Performance, and Early Growth of Hatchling Wall Lizards (Podarcis muralis)

Influence of Incubation Temperature on Morphology, Locomotor Performance, and Early Growth of Hatchling Wall Lizards (Podarcis muralis) JEZ 0774 422 F. BRAÑA JOURNAL AND OF X. JI EXPERIMENTAL ZOOLOGY 286:422 433 (2000) Influence of Incubation Temperature on Morphology, Locomotor Performance, and Early Growth of Hatchling Wall Lizards (Podarcis

More information

Maternally chosen nest sites positively affect multiple components of offspring fitness in a lizard

Maternally chosen nest sites positively affect multiple components of offspring fitness in a lizard Advance Access published August 29, 2012 doi:10.1093/beheco/ars133 Original Article Maternally chosen nest sites positively affect multiple components of offspring fitness in a lizard Aaron M. Reedy, a

More information

EFFECTS OF CROWDING ON REPRODUCTIVE TRAITS OF WESTERN FENCE LIZARDS, SCELOPORUS OCCIDENTALIS

EFFECTS OF CROWDING ON REPRODUCTIVE TRAITS OF WESTERN FENCE LIZARDS, SCELOPORUS OCCIDENTALIS Herpetological Conservation and Biology 8(1):251 257. Submitted: 6 February 2012; Accepted: 8 February 2013; Published: 30 April 2013. EFFECTS OF CROWDING ON REPRODUCTIVE TRAITS OF WESTERN FENCE LIZARDS,

More information

School of Zoology, University of Tasmania, PO Box 252C-05, Tas, 7001, Australia

School of Zoology, University of Tasmania, PO Box 252C-05, Tas, 7001, Australia Functional Ecology 2000 Maternal basking opportunity affects juvenile phenotype Blackwell Science, Ltd in a viviparous lizard E. WAPSTRA School of Zoology, University of Tasmania, PO Box 252C-05, Tas,

More information

Evidence of divergent growth rates among populations of the lizard Anolis carolinensis based on experimental manipulations of egg size

Evidence of divergent growth rates among populations of the lizard Anolis carolinensis based on experimental manipulations of egg size Popul Ecol (2010) 52:113 122 DOI 10.1007/s10144-009-0167-z ORIGINAL ARTICLE Evidence of divergent growth rates among populations of the lizard Anolis carolinensis based on experimental manipulations of

More information

Social and Thermal Cues Influence Nest-site Selection in a Nocturnal Gecko, Oedura lesueurii

Social and Thermal Cues Influence Nest-site Selection in a Nocturnal Gecko, Oedura lesueurii RESEARCH PAPER Social and Thermal Cues Influence Nest-site Selection in a Nocturnal Gecko, Oedura lesueurii David A. Pike*, Jonathan K. Webb* & Robin M. Andrews * School of Biological Sciences A08, University

More information

Short-term Water Potential Fluctuations and Eggs of the Red-eared Slider Turtle (Trachemys scripta elegans)

Short-term Water Potential Fluctuations and Eggs of the Red-eared Slider Turtle (Trachemys scripta elegans) Zoology and Genetics Publications Zoology and Genetics 2001 Short-term Water Potential Fluctuations and Eggs of the Red-eared Slider Turtle (Trachemys scripta elegans) John K. Tucker Illinois Natural History

More information

BULLETIN. Chicago Herpetological Society

BULLETIN. Chicago Herpetological Society BULLETIN of the Chicago Herpetological Society Volume 38, Number 7 July 2003 BULLETIN OF THE CHICAGO HERPETOLOGICAL SOCIETY Volume 38, Number 7 July 2003 Environmental and Maternal Influences on Eggs and

More information

Wen SHEN 1, Jianchi PEI 2, Longhui LIN 3* and Xiang JI Introduction

Wen SHEN 1, Jianchi PEI 2, Longhui LIN 3* and Xiang JI Introduction Asian Herpetological Research 2017, 8(4): 262 268 DOI: 10.16373/j.cnki.ahr.170029 ORIGINAL ARTICLE Effects of Constant versus Fluctuating Incubation Temperatures on Hatching Success, Incubation Length,

More information

A comparison of placental tissue in the skinks Eulamprus tympanum and E. quoyii. Yates, Lauren A.

A comparison of placental tissue in the skinks Eulamprus tympanum and E. quoyii. Yates, Lauren A. A comparison of placental tissue in the skinks Eulamprus tympanum and E. quoyii Yates, Lauren A. Abstract: The species Eulamprus tympanum and Eulamprus quoyii are viviparous skinks that are said to have

More information

Offspring performance and the adaptive benefits of. prolonged pregnancy: experimental tests in a viviparous lizard

Offspring performance and the adaptive benefits of. prolonged pregnancy: experimental tests in a viviparous lizard Functional Ecology 2009, 23, 818 825 doi: 10.1111/j.1365-2435.2009.01544.x Offspring performance and the adaptive benefits of Blackwell Publishing Ltd prolonged pregnancy: experimental tests in a viviparous

More information

THE concept that reptiles have preferred

THE concept that reptiles have preferred Copeia, 2000(3), pp. 841 845 Plasticity in Preferred Body Temperature of Young Snakes in Response to Temperature during Development GABRIEL BLOUIN-DEMERS, KELLEY J. KISSNER, AND PATRICK J. WEATHERHEAD

More information

COMPARING BODY CONDITION ESTIMATES OF ZOO BROTHER S ISLAND TUATARA (SPHENODON GUNTHERI) TO THAT OF THE WILD, A CLINICAL CASE

COMPARING BODY CONDITION ESTIMATES OF ZOO BROTHER S ISLAND TUATARA (SPHENODON GUNTHERI) TO THAT OF THE WILD, A CLINICAL CASE COMPARING BODY CONDITION ESTIMATES OF ZOO BROTHER S ISLAND TUATARA (SPHENODON GUNTHERI) TO THAT OF THE WILD, A CLINICAL CASE Kyle S. Thompson, BS,¹, ²* Michael L. Schlegel, PhD, PAS² ¹Oklahoma State University,

More information

Natural History Note

Natural History Note vol. 176, no. 4 the american naturalist october 2010 Natural History Note The Physiological Basis of Geographic Variation in Rates of Embryonic Development within a Widespread Lizard Species Wei-Guo Du,

More information

Evolution of viviparity in warm-climate lizards: an experimental test of the maternal manipulation hypothesis

Evolution of viviparity in warm-climate lizards: an experimental test of the maternal manipulation hypothesis doi:10.1111/j.1420-9101.2006.01296.x Evolution of viviparity in warm-climate lizards: an experimental test of the maternal manipulation hypothesis X. JI,* C.-X. LIN, à L.-H. LIN,* Q.-B. QIUà &Y.DU à *Jiangsu

More information

University of Canberra. This thesis is available in print format from the University of Canberra Library.

University of Canberra. This thesis is available in print format from the University of Canberra Library. University of Canberra This thesis is available in print format from the University of Canberra Library. If you are the author of this thesis and wish to have the whole thesis loaded here, please contact

More information

A Population Analysis of the Common Wall Lizard Podarcis muralis in Southwestern France

A Population Analysis of the Common Wall Lizard Podarcis muralis in Southwestern France - 513 - Studies in Herpetology, Rocek Z. (ed.) pp. 513-518 Prague 1986 A Population Analysis of the Common Wall Lizard Podarcis muralis in Southwestern France R. BARBAULT and Y. P. MOU Laboratoire d'ecologie

More information

Sex-based hatching asynchrony in an oviparous lizard (Bassiana duperreyi, Scincidae)

Sex-based hatching asynchrony in an oviparous lizard (Bassiana duperreyi, Scincidae) Austral Ecology (2007) 32, 502 508 doi:10.1111/j.1442-9993.2007.01722.x Sex-based hatching asynchrony in an oviparous lizard (Bassiana duperreyi, Scincidae) RAJKUMAR S. RADDER AND RICHARD SHINE* School

More information

Like mother, like daughter: inheritance of nest-site

Like mother, like daughter: inheritance of nest-site Like mother, like daughter: inheritance of nest-site location in snakes Gregory P. Brown and Richard Shine* School of Biological Sciences A0, University of Sydney, NSW 00, Australia *Author for correspondence

More information

DECREASED SPRINT SPEED AS A COST OF REPRODUCTION IN THE LIZARD SCELOPORUS OCCIDENTALS: VARIATION AMONG POPULATIONS

DECREASED SPRINT SPEED AS A COST OF REPRODUCTION IN THE LIZARD SCELOPORUS OCCIDENTALS: VARIATION AMONG POPULATIONS J. exp. Biol. 155, 323-336 (1991) 323 Printed in Great Britain The Company of Biologists Limited 1991 DECREASED SPRINT SPEED AS A COST OF REPRODUCTION IN THE LIZARD SCELOPORUS OCCIDENTALS: VARIATION AMONG

More information

Maturity and Other Reproductive Traits of the Kanahebi Lizard Takydromus tachydromoides (Sauria, Lacertidae) in Mito

Maturity and Other Reproductive Traits of the Kanahebi Lizard Takydromus tachydromoides (Sauria, Lacertidae) in Mito Japanese Journal of Herpetology 9 (2): 46-53. 1981. Maturity and Other Reproductive Traits of the Kanahebi Lizard Takydromus tachydromoides (Sauria, Lacertidae) in Mito Sen TAKENAKA SUMMARY: Reproduction

More information

JEZ Part A: Comparative Experimental Biology. An experimental test of the effects of fluctuating incubation temperatures on hatchling phenotype

JEZ Part A: Comparative Experimental Biology. An experimental test of the effects of fluctuating incubation temperatures on hatchling phenotype An experimental test of the effects of fluctuating incubation temperatures on hatchling phenotype Journal: Manuscript ID: Wiley - Manuscript type: Date Submitted by the Author: JEZ Part A: Physiology and

More information

Lizard malaria: cost to vertebrate host's reproductive success

Lizard malaria: cost to vertebrate host's reproductive success Parasilology (1983), 87, 1-6 1 With 2 figures in the text Lizard malaria: cost to vertebrate host's reproductive success J. J. SCHALL Department of Zoology, University of Vermont, Burlington, Vermont 05405,

More information

The influence of propagule size and maternal nest-site. selection on survival and behaviour of neonate turtles. J. J. KOLBE* and F. J.

The influence of propagule size and maternal nest-site. selection on survival and behaviour of neonate turtles. J. J. KOLBE* and F. J. Functional Ecology 2001 The influence of propagule size and maternal nest-site Blackwell Science Ltd selection on survival and behaviour of neonate turtles J. J. KOLBE* and F. J. JANZEN Department of Zoology

More information

HERPETOLOGICA VOL. 68 JUNE 2012 NO. 2 LIN SCHWARZKOPF 1,3 AND ROBIN M. ANDREWS 2

HERPETOLOGICA VOL. 68 JUNE 2012 NO. 2 LIN SCHWARZKOPF 1,3 AND ROBIN M. ANDREWS 2 HERPETOLOGICA VOL. 68 JUNE 2012 NO. 2 Herpetologica, 68(2), 2012, 147 159 E 2012 by The Herpetologists League, Inc. ARE MOMS MANIPULATIVE OR JUST SELFISH? EVALUATING THE MATERNAL MANIPULATION HYPOTHESIS

More information

Testing the Persistence of Phenotypic Plasticity After Incubation in the Western Fence Lizard, Sceloporus Occidentalis

Testing the Persistence of Phenotypic Plasticity After Incubation in the Western Fence Lizard, Sceloporus Occidentalis Claremont Colleges Scholarship @ Claremont All HMC Faculty Publications and Research HMC Faculty Scholarship 1-1-2007 Testing the Persistence of Phenotypic Plasticity After Incubation in the Western Fence

More information

Abstract. Keywords: Introduction

Abstract. Keywords: Introduction doi: 1.1111/j.14-911.12.2575.x Altitudinal variation in egg retention and rates of embryonic development in oviparous Zootoca vivipara fits predictions from the cold-climate model on the evolution of viviparity

More information

Parthenogenesis in Varanus ornatus, the Ornate Nile Monitor.

Parthenogenesis in Varanus ornatus, the Ornate Nile Monitor. Parthenogenesis in Varanus ornatus, the Ornate Nile Monitor. Parthenogenesis in varanids has been reported in two other species of monitor, the Komodo dragon, Varanus komodiensis (Watts et al) and the

More information

Phenotypic variation in smooth softshell turtles (Apalone mutica) from eggs incubated in constant versus fluctuating temperatures

Phenotypic variation in smooth softshell turtles (Apalone mutica) from eggs incubated in constant versus fluctuating temperatures Oecologia (2003) 134:182 188 DOI 10.1007/s00442-002-1109-z ECOPHYSIOLOGY Grant M. Ashmore Fredric J. Janzen Phenotypic variation in smooth softshell turtles (Apalone mutica) from eggs incubated in constant

More information

Weaver Dunes, Minnesota

Weaver Dunes, Minnesota Hatchling Orientation During Dispersal from Nests Experimental analyses of an early life stage comparing orientation and dispersal patterns of hatchlings that emerge from nests close to and far from wetlands

More information

Supporting Online Material for

Supporting Online Material for www.sciencemag.org/cgi/content/full/314/5802/1111/dc1 Supporting Online Material for Rapid Temporal Reversal in Predator-Driven Natural Selection Jonathan B. Losos,* Thomas W. Schoener, R. Brian Langerhans,

More information

Husbandry and Reproduction of Varanus glauerti in Captivity

Husbandry and Reproduction of Varanus glauerti in Captivity Biawak, 4(3), pp. 103-107 2010 by International Varanid Interest Group Husbandry and Reproduction of Varanus glauerti in Captivity MARTIJN DE ZEEUW Hazerswoude-Dorp, The Netherlands E-mail: Martijn@odatria.nl

More information

Ecological Archives E A2

Ecological Archives E A2 Ecological Archives E089-034-A2 David A. Pike, Ligia Pizzatto, Brian A. Pike, and Richard Shine. 2008. Estimating survival rates of uncatchable animals: the myth high juvenile mortality in reptiles. Ecology

More information

Corn Snake Care Sheet

Corn Snake Care Sheet Corn Snake Care Sheet Temperament With the odd exception, Corn Snakes are calm, docile, placid snakes that are hardy and thrive very well in captivity. Due to their temperament Corn Snakes are a recommended

More information

THE adaptive significance, if any, of temperature-dependent

THE adaptive significance, if any, of temperature-dependent Copeia, 2003(2), pp. 366 372 Nest Temperature Is Not Related to Egg Size in a Turtle with Temperature-Dependent Sex Determination CARRIE L. MORJAN AND FREDRIC J. JANZEN A recent hypothesis posits that

More information

Offspring size number strategies: experimental manipulation of offspring size in a viviparous lizard (Lacerta vivipara)

Offspring size number strategies: experimental manipulation of offspring size in a viviparous lizard (Lacerta vivipara) Functional Ecology 2002 Blackwell Oxford, FEC Functional 0269-8463 British February 16 1000 Ecological UK 2002 Science Ecology Ltd Society, 2002 TECHNICAL REPORT Allometric M. Olsson et engineering al.

More information

Phenotypic Responses of Hatchlings to Constant Versus Fluctuating Incubation Temperatures in the Multi-banded Krait, Bungarus multicintus (Elapidae)

Phenotypic Responses of Hatchlings to Constant Versus Fluctuating Incubation Temperatures in the Multi-banded Krait, Bungarus multicintus (Elapidae) ZOOLOGICAL SCIENCE 24: 384 390 (2007) 2007 Zoological Society of Japan Phenotypic Responses of Hatchlings to Constant Versus Fluctuating Incubation Temperatures in the Multi-banded Krait, Bungarus multicintus

More information

Variation of Chicken Embryo Development by Temperature Influence. Anna Morgan Miller. Rockdale Magnet School for Science and Technology

Variation of Chicken Embryo Development by Temperature Influence. Anna Morgan Miller. Rockdale Magnet School for Science and Technology Variation of Chicken Embryo Development by Temperature Influence Anna Morgan Miller Rockdale Magnet School for Science and Technology Anna Morgan Miller Rockdale Magnet School 1174 Bulldog Circle Conyers,

More information

RESEARCH ARTICLE Potentially adaptive effects of maternal nutrition during gestation on offspring phenotype of a viviparous reptile

RESEARCH ARTICLE Potentially adaptive effects of maternal nutrition during gestation on offspring phenotype of a viviparous reptile 4234 The Journal of Experimental Biology 214, 4234-4239 2011. Published by The Company of Biologists Ltd doi:10.1242/jeb.057349 RESEARCH ARTICLE Potentially adaptive effects of maternal nutrition during

More information

WATER plays an important role in all stages

WATER plays an important role in all stages Copeia, 2002(1), pp. 220 226 Experimental Analysis of an Early Life-History Stage: Water Loss and Migrating Hatchling Turtles JASON J. KOLBE AND FREDRIC J. JANZEN The effect of water dynamics is well known

More information

Chameleons: Biology, Husbandry and Disease Prevention. Paul Stewart, DVM. Origin: Africa (40% of species) and Madagascar (40% of species)

Chameleons: Biology, Husbandry and Disease Prevention. Paul Stewart, DVM. Origin: Africa (40% of species) and Madagascar (40% of species) Chameleons: Biology, Husbandry and Disease Prevention By Paul Stewart, DVM Number of Species: 150 identified Size: From 3.3 cm to 68 cm in length Origin: Africa (40% of species) and Madagascar (40% of

More information

Thermal constraints on embryonic development as a proximate cause for. elevational range limits in two Mediterranean lacertid lizards

Thermal constraints on embryonic development as a proximate cause for. elevational range limits in two Mediterranean lacertid lizards 1 2 3 4 5 Thermal constraints on embryonic development as a proximate cause for elevational range limits in two Mediterranean lacertid lizards 6 7 8 Camila Monasterio 1,3,4, Luke P. Shoo 2,*, Alfredo Salvador

More information

Latent Effects of Egg Incubation Temperature on Growth in the Lizard Anolis carolinensis

Latent Effects of Egg Incubation Temperature on Growth in the Lizard Anolis carolinensis JOURNAL OF EXPERIMENTAL ZOOLOGY 309A (2008) A Journal of Integrative Biology Latent Effects of Egg Incubation Temperature on Growth in the Lizard Anolis carolinensis RACHEL M. GOODMAN Department of Ecology

More information

Who Cares? The Evolution of Parental Care in Squamate Reptiles. Ben Halliwell Geoffrey While, Tobias Uller

Who Cares? The Evolution of Parental Care in Squamate Reptiles. Ben Halliwell Geoffrey While, Tobias Uller Who Cares? The Evolution of Parental Care in Squamate Reptiles Ben Halliwell Geoffrey While, Tobias Uller 1 Parental Care any instance of parental investment that increases the fitness of offspring 2 Parental

More information

A NOVEL PATTERN OF EMBRYONIC NUTRITION IN A VIVIPAROUS REPTILE

A NOVEL PATTERN OF EMBRYONIC NUTRITION IN A VIVIPAROUS REPTILE J. exp. Biol. 174, 97 108 (1993) Printed in Great Britain The Company of Biologists Limited 1993 97 A NOVEL PATTERN OF EMBRYONIC NUTRITION IN A VIVIPAROUS REPTILE BY JAMES R. STEWART AND MICHAEL B. THOMPSON

More information

Sheikh Muhammad Abdur Rashid Population ecology and management of Water Monitors, Varanus salvator (Laurenti 1768) at Sungei Buloh Wetland Reserve,

Sheikh Muhammad Abdur Rashid Population ecology and management of Water Monitors, Varanus salvator (Laurenti 1768) at Sungei Buloh Wetland Reserve, Author Title Institute Sheikh Muhammad Abdur Rashid Population ecology and management of Water Monitors, Varanus salvator (Laurenti 1768) at Sungei Buloh Wetland Reserve, Singapore Thesis (Ph.D.) National

More information

Reproductive physiology and eggs

Reproductive physiology and eggs Reproductive physiology and eggs Class Business Reading for this lecture Required. Gill: Chapter 14 1. Reproductive physiology In lecture I will only have time to go over reproductive physiology briefly,

More information

Climate change impacts on fitness depend on nesting habitat in lizards

Climate change impacts on fitness depend on nesting habitat in lizards Functional Ecology 2011, 25, 1125 1136 doi: 10.1111/j.1365-2435.2011.01855.x Climate change impacts on fitness depend on nesting habitat in lizards Wen-San Huang*,1 and David A. Pike 2 1 Department of

More information

DEVELOPMENTAL SUCCESS, STABILITY, AND PLASTICITY IN CLOSELY RELATED PARTHENOGENETIC AND SEXUAL LIZARDS (HETERONOTIA, GEKKONIDAE)

DEVELOPMENTAL SUCCESS, STABILITY, AND PLASTICITY IN CLOSELY RELATED PARTHENOGENETIC AND SEXUAL LIZARDS (HETERONOTIA, GEKKONIDAE) DEVELOPMENTAL SUCCESS, STABILY, AND PLASTICY IN CLOSELY RELATED PARTHENOGENETIC AND SEXUAL LIZARDS (HETERONOTIA, GEKKONIDAE) Author(s) :Michael Kearney and Richard Shine Source: Evolution, 58(7):560-57.

More information

Sprint speed capacity of two alpine skink species, Eulamprus kosciuskoi and Pseudemoia entrecasteauxii

Sprint speed capacity of two alpine skink species, Eulamprus kosciuskoi and Pseudemoia entrecasteauxii Sprint speed capacity of two alpine skink species, Eulamprus kosciuskoi and Pseudemoia entrecasteauxii Isabella Robinson, Bronte Sinclair, Holly Sargent, Xiaoyun Li Abstract As global average temperatures

More information

A description of an Indo-Chinese rat snake (Ptyas korros [Schlegel, 1837]) clutch, with notes on an instance of twinning

A description of an Indo-Chinese rat snake (Ptyas korros [Schlegel, 1837]) clutch, with notes on an instance of twinning 1 2 A description of an Indo-Chinese rat snake (Ptyas korros [Schlegel, 1837]) clutch, with notes on an instance of twinning 3 4 Simon Dieckmann 1, Gerrut Norval 2 * and Jean-Jay Mao 3 5 6 7 8 9 10 11

More information

The effect of short-term weather fluctuations on temperatures inside lizard nests, and on the phenotypic traits of hatchling lizards

The effect of short-term weather fluctuations on temperatures inside lizard nests, and on the phenotypic traits of hatchling lizards BzoEogical Journal of the Linnean Society (2001), 72: 555565. With 5 figures doi:10.1006/bij1.2000.0516, available online at http;//www.idealibrary.com on I Of @ The effect of shortterm weather fluctuations

More information

LIFE-HISTORY VARIATION IN THE SAGEBRUSH LIZARD: PHENOTYPIC PLASTICITY OR LOCAL ADAPTATION?

LIFE-HISTORY VARIATION IN THE SAGEBRUSH LIZARD: PHENOTYPIC PLASTICITY OR LOCAL ADAPTATION? Ecology, 84(6), 003, pp. 64 634 003 by the Ecological Society of America LIFE-HISTORY VARIATION IN THE SAGEBRUSH LIZARD: PHENOTYPIC PLASTICITY OR LOCAL ADAPTATION? MICHAEL W. SEARS,3 AND MICHAEL J. ANGILLETTA,

More information

Station 1 1. (3 points) Identification: Station 2 6. (3 points) Identification:

Station 1 1. (3 points) Identification: Station 2 6. (3 points) Identification: SOnerd s 2018-2019 Herpetology SSSS Test 1 SOnerd s SSSS 2018-2019 Herpetology Test Station 20 sounds found here: https://drive.google.com/drive/folders/1oqrmspti13qv_ytllk_yy_vrie42isqe?usp=sharing Station

More information

INDIVIDUAL IDENTIFICATION OF GREEN TURTLE (CHELONIA MYDAS) HATCHLINGS

INDIVIDUAL IDENTIFICATION OF GREEN TURTLE (CHELONIA MYDAS) HATCHLINGS INDIVIDUAL IDENTIFICATION OF GREEN TURTLE (CHELONIA MYDAS) HATCHLINGS Ellen Ariel, Loïse Corbrion, Laura Leleu and Jennifer Brand Report No. 15/55 Page i INDIVIDUAL IDENTIFICATION OF GREEN TURTLE (CHELONIA

More information

Calcium provision to oviparous and viviparous embryos of the reproductively bimodal lizard Lacerta (Zootoca) vivipara

Calcium provision to oviparous and viviparous embryos of the reproductively bimodal lizard Lacerta (Zootoca) vivipara 2520 The Journal of Experimental Biology 212, 2520-2524 Published by The Company of Biologists 2009 doi:10.1242/jeb.030643 Calcium provision to oviparous and viviparous embryos of the reproductively bimodal

More information

Geographical differences in maternal basking behaviour and offspring growth rate in a climatically widespread viviparous reptile

Geographical differences in maternal basking behaviour and offspring growth rate in a climatically widespread viviparous reptile 2014. Published by The Company of Biologists Ltd (2014) 217, 1175-1179 doi:10.1242/jeb.089953 RESEARCH ARTICLE Geographical differences in maternal basking behaviour and offspring growth rate in a climatically

More information

Nest Site Creation and Maintenance as an Effective Tool in Species Recovery

Nest Site Creation and Maintenance as an Effective Tool in Species Recovery Nest Site Creation and Maintenance as an Effective Tool in Species Recovery Scott D. Gillingwater Species At Risk Biologist Upper Thames River Conservation Authority Where and Why? The successful creation

More information

NOTES ON THE ECOLOGY AND NATURAL HISTORY OF TWO SPECIES OF EGERNIA (SCINCIDAE) IN WESTERN AUSTRALIA

NOTES ON THE ECOLOGY AND NATURAL HISTORY OF TWO SPECIES OF EGERNIA (SCINCIDAE) IN WESTERN AUSTRALIA NOTES ON THE ECOLOGY AND NATURAL HISTORY OF TWO SPECIES OF EGERNIA (SCINCIDAE) IN WESTERN AUSTRALIA By ERIC R. PIANKA Integrative Biology University of Texas at Austin Austin, Texas 78712 USA Email: erp@austin.utexas.edu

More information

Ambystoma jeffersonianum at Tuscarora Pond, Douthat State Park: Data from a population at the Southern extent of the species range

Ambystoma jeffersonianum at Tuscarora Pond, Douthat State Park: Data from a population at the Southern extent of the species range Ambystoma jeffersonianum at Tuscarora Pond, Douthat State Park: Data from a population at the Southern extent of the species range Paul Sattler Jason Gibson Biology Department Danville City Schools Liberty

More information

The sooner the better: reproductive phenology drives ontogenetic trajectories in a temperate squamate (Podarcis muralis)

The sooner the better: reproductive phenology drives ontogenetic trajectories in a temperate squamate (Podarcis muralis) bs_bs_banner Biological Journal of the Linnean Society, 2013, 108, 384 395. With 5 figures The sooner the better: reproductive phenology drives ontogenetic trajectories in a temperate squamate (Podarcis

More information

Phenotypic and fitness consequences of maternal nest-site choice across multiple early life stages

Phenotypic and fitness consequences of maternal nest-site choice across multiple early life stages Ecology, Evolution and Organismal Biology Publications Ecology, Evolution and Organismal Biology 2-2013 Phenotypic and fitness consequences of maternal nest-site choice across multiple early life stages

More information

Phenotypic and fitness consequences of maternal nest-site choice across multiple early life stages

Phenotypic and fitness consequences of maternal nest-site choice across multiple early life stages Ecology, 94(2), 2013, pp. 336 345 Ó 2013 by the Ecological Society of America Phenotypic and fitness consequences of maternal nest-site choice across multiple early life stages TIMOTHY S. MITCHELL, 1 DANIEL

More information

A Rhode Island Non-Profit Organization Bearded Dragon Care

A Rhode Island Non-Profit Organization Bearded Dragon Care www.rirescue.org A Rhode Island Non-Profit Organization Bearded Dragon Care Bearded dragons are solitary lizards of the Agama family who originate from the grasslands of Australia and spend most of their

More information

Rigid Shells Enhance Survival of Gekkotan Eggs

Rigid Shells Enhance Survival of Gekkotan Eggs RESEARCH ARTICLE Rigid Shells Enhance Survival of Gekkotan Eggs ROBIN M. ANDREWS* Department of Biological Sciences, Virginia Tech, Blacksburg, Virginia ABSTRACT 323A:607 615, 2015 The majority of lizards

More information

and hydration of hatchling Painted Turtles, Chrysemys picta

and hydration of hatchling Painted Turtles, Chrysemys picta Functional Ecology 21 Environmentally induced variation in size, energy reserves Blackwell Science, Ltd and hydration of hatchling Painted Turtles, Chrysemys picta G. C. PACKARD and M. J. PACKARD Colorado

More information

The effect of weaning weight on subsequent lamb growth rates

The effect of weaning weight on subsequent lamb growth rates Proceedings of the New Zealand Grassland Association 62: 75 79 (2000) 75 The effect of weaning weight on subsequent lamb growth rates T.J. FRASER and D.J. SAVILLE AgResearch, PO Box 60, Lincoln, Canterbury

More information

Thermal and fitness-related consequences of nest location in Painted Turtles (Chrysemys picta)

Thermal and fitness-related consequences of nest location in Painted Turtles (Chrysemys picta) Functional Ecology 1999 ORIGINAL ARTICLE OA 000 EN Thermal and fitness-related consequences of nest location in Painted Turtles (Chrysemys picta) D. W. WEISROCK and F. J. JANZEN* Department of Zoology

More information

SUMMARY OF THESIS. Chapter VIII "The place of research, its purpose, the biological material and method"

SUMMARY OF THESIS. Chapter VIII The place of research, its purpose, the biological material and method SUMMARY OF THESIS Raising Japanese quail is a global activity still limited compared with growth of hens and broilers, but with great prospects for the development of characteristics and adaptability of

More information

Keywords Geographic variation Lizards Reproductive output Reproductive mode Maternal body size Offspring size RESEARCH ARTICLE

Keywords Geographic variation Lizards Reproductive output Reproductive mode Maternal body size Offspring size RESEARCH ARTICLE Evol Biol (2013) 40:420 438 DOI 10.1007/s11692-013-9247-2 RESEARCH ARTICLE Variation of Reproductive Traits and Female Body Size in the Most Widely-Ranging Terrestrial Reptile: Testing the Effects of Reproductive

More information

Maternal Effects in the Green Turtle (Chelonia mydas)

Maternal Effects in the Green Turtle (Chelonia mydas) Maternal Effects in the Green Turtle (Chelonia mydas) SUBMITTED BY SAM B. WEBER TO THE UNIVERSITY OF EXETER AS A THESIS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN BIOLOGY; 8 TH JUNE 2010 This thesis is

More information

Effect of Tail Loss on Sprint Speed and Growth in Newborn Skinks, Niveoscincus metallicus

Effect of Tail Loss on Sprint Speed and Growth in Newborn Skinks, Niveoscincus metallicus Effect of Tail Loss on Sprint Speed and Growth in Newborn Skinks, Niveoscincus metallicus Author(s) :David G. Chapple, Colin J. McCoull, Roy Swain Source: Journal of Herpetology, 38(1):137-140. 2004. Published

More information

Modelling exposure to selected temperature during pregnancy: the limitations of squamate viviparity in a cool-climate environment

Modelling exposure to selected temperature during pregnancy: the limitations of squamate viviparity in a cool-climate environment Biological Journal of the Linnean Society, 2009, 96, 541 552. With 6 figures Modelling exposure to selected temperature during pregnancy: the limitations of squamate viviparity in a cool-climate environment

More information

Effects of Thermal and Hydric Conditions on Egg Incubation and Hatchling Phenotypes in Two Phrynocephalus Lizards

Effects of Thermal and Hydric Conditions on Egg Incubation and Hatchling Phenotypes in Two Phrynocephalus Lizards Asian Herpetological Research 2012, 3(3): 184 191 DOI: 10.3724/SP.J.1245.2012.00184 Effects of Thermal and Hydric Conditions on Egg Incubation and Hatchling Phenotypes in Two Phrynocephalus Lizards Xiaolong

More information

These small issues are easily addressed by small changes in wording, and should in no way delay publication of this first- rate paper.

These small issues are easily addressed by small changes in wording, and should in no way delay publication of this first- rate paper. Reviewers' comments: Reviewer #1 (Remarks to the Author): This paper reports on a highly significant discovery and associated analysis that are likely to be of broad interest to the scientific community.

More information

LETTER TO THE EDITOR. The Evolution of Viviparity and Placentation Revisted

LETTER TO THE EDITOR. The Evolution of Viviparity and Placentation Revisted J. theor. Biol. (1997) 185, 129 135 The Evolution of Viviparity and Placentation Revisted Most studies on the evolution of viviparity in squamate reptiles have focused on one or more of the following questions:

More information

Temperature-Dependent Sex Determination in the Leopard Gecko, Eublepharis macularius

Temperature-Dependent Sex Determination in the Leopard Gecko, Eublepharis macularius THE JOURNAL OF EXPERIMENTAL ZOOLOGY 265579-683 (1993) RAPID COMMUNICATION Temperature-Dependent Sex Determination in the Leopard Gecko, Eublepharis macularius BRIAN E. VIETS, ALAN TOUSIGNANT, MICHAEL A.

More information

Viviparity in high altitude Phrynocephalus lizards is adaptive because embryos cannot fully develop without maternal thermoregulation

Viviparity in high altitude Phrynocephalus lizards is adaptive because embryos cannot fully develop without maternal thermoregulation DOI 10.1007/s00442-013-2811-8 Physiological ecology - Original research Viviparity in high altitude Phrynocephalus lizards is adaptive because embryos cannot fully develop without maternal thermoregulation

More information

Do operational sex ratios influence sex allocation in viviparous lizards with temperature-dependent sex determination?

Do operational sex ratios influence sex allocation in viviparous lizards with temperature-dependent sex determination? doi:10.1111/j.1420-9101.2006.01086.x Do operational sex ratios influence sex allocation in viviparous lizards with temperature-dependent sex determination? D. J. ALLSOP, D. A. WARNER, T. LANGKILDE, 1 W.

More information

D. Burke \ Oceans First, Issue 3, 2016, pgs

D. Burke \ Oceans First, Issue 3, 2016, pgs Beach Shading: A tool to mitigate the effects of climate change on sea turtles Daniel Burke, Undergraduate Student, Dalhousie University Abstract Climate change may greatly impact sea turtles as rising

More information